REVIEW ARTICLE

Reproductive strategies in the lizard,

Calotes versicolor
Bhagyashri A. Shanbhag
Department of Zoology, Karnatak University, Dharwad 580 003, India

Animal species have evolved diverse reproductive

strategies to maximize their reproductive success. Studies on the Indian garden lizard (Calotes versicolor)
have recently revealed many fascinating reproductive
strategies such as manipulation of clutch and egg size
depending upon breeding timing, in spite of pelvic
aperture constraint; switching from capital breeder to
income breeder; female sperm storage as long as six
months; oviductal egg retention for six months or
even more under adverse conditions, by manifesting
embryonic diapause and preventing oviposition. The
findings on C. versicolor provide clues to the evolution
of viviparity. In addition, perception of socio-sexual
(tactile, visual and chemical) cues from conspecifics of
opposite sex plays an important role in their own
gonadal recrudescence and reproduction. The significance and mechanisms governing the evolution of diverse reproductive strategies in the lizard are discussed.
THE garden lizard, Calotes versicolor (family Agamidae)
is widely distributed in India and it can be maintained
with ease in captivity. It is a multi-clutched, seasonal
breeder with long breeding season (May to October)1,2. It
exhibits polyautochrony, i.e. ovulates many eggs simultaneously from both the ovaries3,4. Information on ovarian follicular kinetics3, plasma sex steroid hormone
profiles during gonadal cycles4, fat body cycles5,6, endocrine regulation of gonads711 and accessory reproductive
organs1214, and role of proximate factors in controlling
gonadal cycles2 is available for the species. In addition,
gonadal sex differentiation15,16 and mechanism controlling the phenomenon1720 are studied in this species. In
recent years, the evolution of reproductive strategies of
C. versicolor has been elucidated. This paper reviews
reproductive strategies like manipulation of fecundity,
switching from capital to income breeder, prolonged egg
retention and embryonic diapause, female sperm-storage
phenomenon, and role of socio-sexual factors in gonadal
development in C. versicolor.

Manipulation of fecundity
The egg-laying pattern in squamates varies. Some species
lay fixed number of eggs in a clutch (determinate clutch
or fixed clutch). For example, most geckos have a clutch
e-mail: bhagyashri_shanbhag@hotmail.com
646

size of two eggs. In contrast, others lay variable number

of eggs (e.g. C. versicolor, Mabuya carinata) in a clutch
(variable clutch). In lizards with fixed as well as with
variable clutch, especially from the tropical region, clutch
frequency may vary with proximate climatic factors and
food availability. On the other hand, in individuals with
variate clutch pattern, both clutch size and frequency may
vary in relation to proximate climatic factors, food availability, fat reserves of the body and maternal body
size21,22. Several studies, especially on chelonians and
squamates, describe how optimal reproductive fitness is
achieved through manipulation of clutch and/or egg
size2123. There are species in which the egg size is optimized in a population irrespective of the body size of the
female24. A fixity in pelvic aperture size leads to optimization of the egg size. However, in some species of
turtles and lizards24,25, the egg size is not optimized and
therefore varies with the body size of the female. It has
been shown in these species that an increase in the body
size is accompanied by a corresponding increase in the
pelvic aperture. Hence, in these cases optimization of egg
size is not obligatory.
The recent studies by Shanbhag et al.26 on fecundity
manipulation by C. versicolor have revealed some interesting phenomena. They showed that the clutch size and
clutch mass exhibit a positive correlation with body size
(snout-vent length, SVL; Figure 1), indicating that reproductive investment is influenced by body size of the
female. Further, the clutch and egg sizes are inversely
related (Figure 2). Thus, a trade-off between clutch size
and egg size (mass and volume) is obvious in C. versicolor and there is no optimization of egg size. More
interestingly, the above study also showed that the total
energy allocated for reproduction (clutch mass) remains
the same in early, mid and late breeding individuals
(Figure 3), even though the fecundity (clutch size) declines in late breeders. Apparently, the energy allocated
for reproduction in each bout of egg production in
C. versicolor is optimized by natural selection provided
other conditions such as food availability, etc. are optimal,
and predator pressure is minimal. Further, in this lizard
SVL is the prime factor determining the clutch mass26.
In C. versicolor, width of the pelvic aperture remains
virtually constant over the entire range of body sizes
after attainment of sexual maturity. Consequently, pelvic
aperture size becomes a constraint and optimization of
eggs become obligatory. If not, a female producing larger
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eggs may become egg-bound due to pelvic constraint and
eventually die. In C. versicolor, late breeders produce
large-sized eggs irrespective of their body size (Figure
3). Thus, producing larger eggs by late breeders could
result in a setback. Yet, these lizards produce eggs of
variable sizes. This intriguing problem was resolved by
carefully analysing the data on egg dimensions21. The
analysis showed that when the lizards produce larger
eggs, they change the egg length rather than the width
(Table 1) over the entire range of SVL to overcome pelvic constraint.
Like in most oviparous vertebrates, female lizards deposit extra yolk in the egg, beyond what is absolutely
necessary for the development of embryo until the hatching stage. Interestingly, though there is a lot of variation
in egg size with respect to early, mid and late breeding in
the season, there is no appreciable difference in size
(SVL) of hatchlings. However, larger eggs of the late
breeding season produce heavier hatchlings and also possess more internalized yolk than those born early in the
breeding season (MayJune)27. Radder et al.27 opine that
producing heavier hatchlings with more internalized yolk
at the end of the breeding season may enhance fitness
of the hatchlings, since late born have to compete for
Figure 1. Positive relationship between body size (SVL) and clutch
size and mass in C. versicolor.

Figure 2. Clutch and egg sizes (mass and volume) and egg length
and width in C. versicolor. Note that clutch size is negatively correlated with egg size and egg length.
CURRENT SCIENCE, VOL. 84, NO. 5, 10 MARCH 2003

Figure 3. Change in clutch size, clutch mass and egg volume in C.

versicolor of different SVL groups with respect to early, mid and late
clutches. Asterisks above standard error bars indicate a significant
difference within the same SVL group among early, mid and late
clutches. Numbers above SE bars indicate sample size. (Adapted from
Shanbhag et al.26).
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Table 1.

Length (L) and width (W) in mm of eggs in Calotes versicolor with respect to SVL
Clutch

SVL (cm)

Early (MayJune)

Mid (JulyAugust)

Late (SeptemberOctober)

< 9.5

L
W

13.18 0.13
7.18 0.18
(8)

12.66 0.21
7.16 0.30
(6)

14.75 0.53*
7.60 0.24
(10)

9.511.0

L
W

12.98 0.17
7.39 0.10
(37)

12.72 0.18
7.22 0.17
(22)

13.75 0.23*
7.70 0.16
(20)

> 11.0

L
W

12.81 0.33
7.31 0.13
(8)

12.93 0.07
7.29 0.38
(7)

14.00 0.42*
7.81 0.20
(8)

*Significant difference within the same SVL group among early, mid and late clutches at
P < 0.05 based on one-way ANOVA using log-transformed data. Numbers in parentheses indicate sample size. (Adapted from Shanbhag et al.26).

food and other resources with older individuals of early

clutches.

Switching from capital to income breeding

All organisms must use their energy resources judicially
to ensure growth, survival and reproduction. An organism
may overcome increased demand of resources/energy for
reproduction by increasing its food intake. The reproductive fitness could be achieved by two ways28. The socalled capital breeders acquire food in advance and
store it endogenously or exogenously until reproduction.
Others, especially those living in a predictable environment with unlimited food resource, adjust their food intake
concurrently with breeding activity (income breeders).
Several species of squamates are known to store energy
in the form of abdominal fat bodies or fat pads as reserve
energy. The energy stores are mainly utilized as reserve
food during hibernation in temperate species. Among
tropical species, the fat bodies play a main supporting
role in reproduction, especially during periods of unpredictable or fluctuating resource availability.
Shanbhag and Prasad5 showed that in C. versicolor, the
abdominal fat bodies have a supportive role in reproduction. An inverse relationship between the fat body mass
and ovarian cycle is found in C. versicolor (Figure 4).
They also demonstrated by experimental manipulations,
such as fatectomy (FBX), ovariectomy (OvX) and E2
treatment to the OvX lizards, that energy stores in the fat
bodies are indeed used for growth of vitellogenic follicles. It is interesting to note that in C. versicolor, for the
vitellogenic growth of the first clutch of eggs, the energy
stores in the form of fat bodies are used up while the
energy requirement for growth of vitellogenic follicles of
subsequent clutches is met through intake of food. Thus,
C. versicolor adopts a strategy of capital breeder for production of the first clutch of the season and later switches
to income breeding by relying upon daily food intake for
sustaining development of subsequent clutches of the
season. Incidentally, food is generally abundant in the
later part of the breeding season5.
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Figure 4. Fat body-somatic index (FBSI) during annual breeding

cycle in C. versicolor. Note that FBSI drops at the formation of first
clutch of the breeding season (arrow) and remains almost nil during the
formation of subsequent clutches.

Sperm storage in females

Several species of reptiles are known to store sperm in the
oviduct for variable length of time depending upon species2. This phenomenon is found among temperate species
in which the gonadal cycles are temporally dissociated.
For instance, the male skink (Hemiergis peronii) inhabiting the temperate region produces sperm in autumn and
also mates with females prior to ovarian recrudescence29.
The testes then regress. The females store sperm in the
oviduct. After emergence from hibernation, females ovulate and use the stored sperm for fertilization. The young
are born in early spring. Sperm storage in females is
obligatory in such cases. The lizard, C. versicolor does
not hibernate in south India. Also, the gonadal cycle in
both the sexes is not temporally separated. Therefore,
sperm storage by females is not obligatory in C. versicolor. Yet, they store sperms in sperm receptacles formed
by the mucosal folds of the oviduct at the uterovaginal
junction30. Survival of the sperm for as long as six
months in the oviduct is an intriguing phenomenon. The
physiological and biochemical factors responsible for
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stood. Sperms in the sperm receptacle are found with
PAS-positive secretion, possibly derived from the secretion of male reproductive tract31. Also, secretions present
in the sperm receptacle in C. versicolor exhibit elevated
activity of acid phosphatase. This enzyme seems to be
derived from the male and principally from the renal sexual segment, and may have a role in sperm sustenance in
the female reproductive tract31. Shanbhag and Prasad32
have experimentally shown the fertilizing ability of
stored sperm in the species. Vitellogenic females reared
in the laboratory in the absence of males were induced to
ovulate by injecting PMSG. The eggs were not only fertilized using stored oviductal sperm, but they also successfully continued embryonic development. The oviductal
sperm storage mechanism is also seen in Psammophilus
dorasalis that lays more than one clutch per breeding
season33. However, oviductal sperm storage is not observed in M. carinata that lays only one clutch during the
breeding season31. This suggests that oviductal sperm
storage strategy has been evolved in tropical lizards that
lay eggs in multiple clutches possibly to eliminate repeated mating and reduce risk of predation. In C. versicolor,
testicular regression commences earlier than that of the
ovaries during a breeding season. In spite of early testicular regression in males, females produce eggs that seem
to be fertilized by sperm stored in the oviduct.

prolonged oviductal egg retention phenomenon when

oviposition sites or certain proximate factors (e.g. severe
cold, lack of rains, etc.) are not congenial34. The egg retention phenomenon was systematically studied under
laboratory condition in C. versicolor by Shanbhag and
co-workers. Normally, the lizards retain eggs in the oviduct for about two weeks before oviposition, when
embryos are around stage 2728 of development32,35.
Radder et al.36 have shown that C. versicolor can retain
eggs in the oviduct (Figure 5) for at least six months
(maximum for any lizard species recorded so far), or
even more in captive specimens. The eggs, if stripped
from the oviduct, produce normal hatchlings following
incubation at ambient temperature. Prolonged egg retention poses at least two problems to the mother. First, the
growing embryos demand more oxygen and moisture.
Secondly, the water imbibed through the parchment-like
shells swells up the eggs and may cause difficulty in their
exit through the pelvic aperture, the size of which is fixed
in C. versicolor. Thus, the pelvic constraint must discourage any increase in egg width. However, C. versicolor has successfully overcome the problems associated
with egg retention for prolonged period by evolving two
mechanisms arresting the embryonic development at
stage 34 (embryonic diapause), and preventing oviductal
contraction, vis--vis expulsion of eggs.

Oviductal egg retention and embryonic diapause

Control of embryonic diapause

Evolution of transition in reproductive mode from egglaying to live-bearing is obvious in all classes of vertebrates, except in the aves. The reptiles are the first vertebrates to conquer life on land by evolving cleidoic eggs
and internal fertilization. Retention of eggs in the oviduct
for short duration (12 weeks) may help completion of
early embryonic development. However, some species of
lizards from temperate as well as tropical regions exhibit

Like in many other squamate species with flexible eggshell, eggs of C. versicolor increase in volume with
embryonic growth following oviposition37. The increase
in size of eggs is mainly due to imbibed moisture. Arresting embryonic growth at stage 34 limits the egg size such

Figure 5. C. versicolor with eggs retained in the oviduct for about 45

days. Arrows show bulges in the lateral abodominal wall due to presence of retained eggs.

Figure 6. Ambient temperature and body temperature of males, nongravid and gravid females. Note that there is a drastic drop in body
temperature of gravid females with prolonged egg retention.

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that it will facilitate their passage through the pelvic
aperture. Further, arresting embryonic growth also reduces physiological demands of otherwise growing embryo
for oxygen and moisture. Shanbhag et al.38 have shown
that the embryonic diapause is brought about in C. versicolor during prolonged egg retention by drastically lowering its body temperature by 35C (Figure 6) throughout
the period. Body temperature of the females with prolonged egg retention hovers around 23C, and thereby
brings about embryonic diapause. In conformity with this
observation is the fact that in vitro embryonic growth in
C. versicolor arrests around 23C (ref. 37).

Endocrine control of prolonged egg retention

Like in other oviparous lizards39,40, in C. versicolor, corpora lutea (CL) are the source of progesterone (P) in
gravid individuals41. However, Shanbhag et al.41 have
shown that in C. versicolor, plasma P levels fall after
mid-gestation, coinciding with trace 3-hydroxysteroid
dehydrogenase (3-HSDH) enzyme activity in CL. These
findings indicate that high levels of P are not essential
after mid-gestation (i.e. after one week) in the lizard.
Interestingly, in lizards with prolonged egg retention, i.e.
beyond two weeks, the plasma P levels rise (Figure 7)
despite the collapse of CL. Interestingly, a marked rise in
3-HSDH activity is then observed in the adrenal glands,
while CL continue to degenerate. Hence, in such lizards
the source of elevated plasma P is the adrenal gland and
not the ovaries41. Thus, in response to egg-retention stress,
the adrenal glands (the main targets of stress) of gravid
lizard secrete high amount of P. Shine and Guillette42 had
hypothesized that prolonged oviductal egg retention, one
of the steps in the evolution of viviparity in reptiles,
might have evolved under stressful ecological conditions,
and hormones from adrenal glands might help in the

Figure 7. Plasma progesterone (P) levels in C. versicolor during

normal gestation and prolonged egg retention period. IIV represent
stages during normal gestation: I, Following ovulation; II,
During egg-shell formation; III, Embryo at stage 1416; IV, Just before
oviposition, stage 2526 embryo. Note that plasma P levels drop and
rise at stage II, drop to basal level prior to oviposition but rise again in
lizards with retained eggs. (Adapted from Shanbhag et al.41).
650

process of egg retention. The findings on C. versicolor,

besides showing that P secreted by the adrenal gland
promotes oviductal egg retention (by its antimyometrial
action), also provide experimental support to the stressinduced evolution of viviparity hypothesis in reptiles.

Role of socio-sexual cues in reproduction

In seasonally breeding reptiles, a gonadal cycle typically
involves recrudescence, breeding and post-breeding quiescence phases. Interaction between external (physical,
climatic and social environment) and internal (neuroendocrine) factors is essential for successful reproduction.
The control of gonadal cycles through endocrine mechanisms and proximate factors is reasonably well-understood for lizards2,43. Little is known about the role of
social cues governing reproduction in lizards. Recent
researches, however, show that social cues are also
among important regulators of reptilian reproduction.
The integration of social cues in lizards may be mediated
through eyes, olfaction and vomeronasal systems with
specific input to specific brain nuclei. In the brain, specific nuclei in the limbic system that concentrate steroid
hormones receive input from these sensory areas, and
these in turn project to the hypothalamus where both
internal and external cues are integrated43. Studies dealing with the role of socio-sexual factors in reptilian
reproduction are mostly carried out on the genus Anolis.
These in general show that among lizards maintained in
groups in the laboratory, the sexual composition of conspecifics of opposite sex, social experience and dominancesubordinance hierarchy through neuroendocrine
mechanisms influence gonadal recrudescence and mating
behaviour of the cagemates4446.
Through elegantly designed experiments, Shanbhag
et al.47 have demonstrated the importance of socio-sexual
factors in gonadal recrudescence and plasma steroid hormone levels in C. versicolor. In this study, adult lizards
were maintained in groups as (1) males and females, (2)
males and females separated by a wire mesh, (3) same
sex groups of males or females, (4) castrated males with
intact females, and (5) OvX females with intact males
from post-breeding to breeding phase. The study has
shown that isolation of lizards from the opposite sex
adversely affects gonadal recrudescence. In all-male
groups, spermatogenetic recrudescence was not observed
(Figure 8 a). The presence of females (intact or ovariectomized) is needed for initiation of spermatogenesis.
Further, males that could view females with intact recrudescing ovaries through a wire mesh completed only one
wave of spermatogenesis. But for sustenance of spermatogenesis, the presence of females with recrudescing
ovaries producing estrogen is needed. Possibly, elevated
E2 levels are needed for production of pheromonal
signals in the females. Therefore, perception of chemical,
tactile and visual cues from the female by the male seems
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wCaM

Figure 9. Plasma T (male) and E2 (female) values in C. versicolor

subjected to various socio-sexual situations. Values are mean SE
(n = 5 for each group). iC, Initial control; aM, All males; aF, All
females; sW, Separated from opposite sex by wire mesh; wCaM, Kept
with castrated males; wOvX, Maintained with OvX females; wM/wF,
Kept with intact males/females; OvX, Ovariectomized females; CaM,
Castrated males; Wild, Wild caught male/female. Non-identical alphabets above the bar indicate significant difference among groups.
(Adapted from Shanbhag et al.47).

Table 2. Effect of group composition on number of vitellogenic follicles and range of largest follicular diameter in female C. versicolor
Group

Figure 8. Transverse sections of the testes. a, Small seminiferous

tubules lined by a few spermatogonia and Sertoli cells in a male from
an all-male group; b, Enlarged seminiferous tubules and all stages of
spermatogenesis, including sperm in lizards housed with intact females.
Scale line indicates 50 m.

to play an important role in sustaining qualitative and

quantitative aspects of spermatogenetic activity (Figure
8 b) and high plasma T levels (Figure 9) in C. versicolor.
Similarly, the presence of intact males with active
spermatogenesis and elevated plasma T level facilitates
recruitment and development of greater number of vitellogenic follicles (Table 2) and elevated plasma E2 levels
(Figure 9) in C. versicolor. The absence of males inhibits
ovarian recrudescence. Females maintained with castrated males or receiving only visual/chemical cue from the
males were unable to exhibit optimum ovarian activity.
Thus, the perception of socio-sexual (tactile, visual and
chemical) cues from the members of the opposite sex is
important for regulating its own gonadal recrudescence
vis--vis reproductive effort in C. versicolor. After all,
the production of gametes is an expensive phenomenon
and their production in the absence of mates in a seasonally breeding species would be a wasteful process.
CURRENT SCIENCE, VOL. 84, NO. 5, 10 MARCH 2003

Initial control
With intact male
With castrated male
Separated by wire mesh
All females
Wild caught*

No. of vitellogenic
follicles

Largest follicular
diameter (mm)

12.20 0.37a
9.20 0.58b
6.80 0.58c

10.60 0.93a

0.590.80a
3.807.96b
3.175.41c
2.783.92c
1.002.29d
2.407.52b

Non-identical superscripts indicate a significant difference between

groups at P < 0.05 analysed using one-way ANOVA. *Wild caught
lizards at termination of the experiment.

Conclusions
Recent researches have shown that the garden lizard
C. versicolor has evolved several strategies to optimize
its reproductive fitness. For instance, plasticity to manipulate clutch and egg size depending upon the breeding
timing is an interesting phenomenon exhibited by it. It
would be interesting to know the pattern of clutch and
egg-size modifications in other agamids and also in other
sympatric lizards occupying different microhabitats.
Female sperm storage and maintenance of viable sperm
for over six months is another important strategy that
seems to have co-evolved with multiple clutch pattern in
C. versicolor. However, the mechanisms for survival of
sperm in the oviduct for a long period are not yet fully
understood and need investigation. DNA fingerprinting
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of stored sperm is needed to resolve whether C. versicolor follows monoandry or polyandry. Prolonged egg
retention in the oviduct is a noteworthy strategy exhibited
by C. versicolor. Whether lizards other than agamids
found in India exhibit such a phenomenon, is unknown.
Exhaustive studies involving more representative species
are needed to generalize the mechanisms involved in prolonged oviductal egg retention in lizards. It should be
noted that prolonged egg retention is a stage in the evolution of viviparity. Studies on C. versicolor show that
such a phenomenon is related to stress (captivity, absence
of suitable conditions for oviposition, and so on) and that
adrenal glands, through secretion of P, promote egg
retention. These findings provide experimental proof to
the stress-induced evolution of viviparity hypothesis.
The most neglected area of research pertains to the role
of social environment and interaction with conspecifics
in the control of gonadal recrudescence in lizards. However, recent studies on C. versicolor show that conspecifics of opposite sex mutually regulate or affect gonadal
recrudescence. Hence, social isolation leads to failure of
gonadal recrudescence and thereby reproduction. Studies
on C. versicolor have thus revealed conclusively the
importance of cues (visual, olfactory and tactile) from
conspecifics of the opposite sex in its own gonadal recrudescence.
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ACKNOWLEDGEMENTS. I thank Prof. S. K. Saidapur for his
valuable suggestions and encouragement. Financial support by a grant
from Department of Science and Technology, New Delhi and UGC
SAP II, New Delhi is acknowledged.
Received 12 August 2002; revised accepted 11 November 2002
CURRENT SCIENCE, VOL. 84, NO. 5, 10 MARCH 2003